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Intensifying wildfire activity and climate change can drive rapid forest compositional shifts. In boreal North America, black spruce shapes forest flammability and depends on fire for regeneration. This relationship has helped black spruce maintain its dominance through much of the Holocene. However, with climate change and more frequent and severe fires, shifts away from black spruce dominance to broadleaf or pine species are emerging, with implications for ecosystem functions including carbon sequestration, water and energy fluxes, and wildlife habitat. Here, we predict that such reductions in black spruce after fire may already be widespread given current trends in climate and fire. To test this, we synthesize data from 1,538 field sites across boreal North America to evaluate compositional changes in tree species following 58 recent fires (1989 to 2014). While black spruce was resilient following most fires (62%), loss of resilience was common, and spruce regeneration failed completely in 18% of 1,140 black spruce sites. In contrast, postfire regeneration never failed in forests dominated by jack pine, which also possesses an aerial seed bank, or broad-leaved trees. More complete combustion of the soil organic layer, which often occurs in better-drained landscape positions and in dryer duff, promoted compositional changes throughout boreal North America. Forests in western North America, however, were more vulnerable to change due to greater long-term climate moisture deficits. While we find considerable remaining resilience in black spruce forests, predicted increases in climate moisture deficits and fire activity will erode this resilience, pushing the system toward a tipping point that has not been crossed in several thousand years.

Changing disturbance regimes and climate can overcome forest ecosystem resilience. Following high-severity fire, forest recovery may be compromised by lack of tree seed sources, warmer and drier postfire climate, or short-interval reburning. A potential outcome of the loss of resilience is the conversion of the prefire forest to a different forest type or nonforest vegetation. Conversion implies major, extensive, and enduring changes in dominant species, life forms, or functions, with impacts on ecosystem services. In the present article, we synthesize a growing body of evidence of fire-driven conversion and our understanding of its causes across western North America. We assess our capacity to predict conversion and highlight important uncertainties. Increasing forest vulnerability to changing fire activity and climate compels shifts in management approaches, and we propose key themes for applied research coproduced by scientists and managers to support decision-making in an era when the prefire forest may not return.

Forest ecosystems are strongly impacted by continuing climate change and increasing disturbance activity, but how forest dynamics will respond remains highly uncertain. Here, we argue that a short time window after disturbance (i.e., a discrete event that disrupts prevailing ecosystem structure and composition and releases resources) is pivotal for future forest development. Trees that establish during this reorganization phase can shape forest structure and composition for centuries, providing operational early indications of forest change. While forest change has been fruitfully studied through a lens of resilience, profound ecological changes can be masked by a resilience versus regime shift dichotomy. We present a framework for characterizing the full spectrum of change after disturbance, analyzing forest reorganization along dimensions of forest structure (number, size, and spatial arrangement of trees) and composition (identity and diversity of tree species). We propose four major pathways through which forest cover can persist but reorganize following disturbance: resilience (no change in structure and composition), restructuring (structure changes but composition does not), reassembly (composition changes but structure does not), and replacement (structure and composition both change). Regime shifts occur when vegetation structure and composition are altered so profoundly that the emerging trajectory leads to nonforest. We identify fundamental processes underpinning forest reorganization which, if disrupted, deflect ecosystems away from resilience. To understand and predict forest reorganization, assessing these processes and the traits modulating them is crucial. A new wave of experiments, measurements, and models emphasizing the reorganization phase will further the capacity to anticipate future forest dynamics.

CONTEXT: An increase in the incidence of large wildfires worldwide has prompted concerns about the resilience of forest ecosystems, particularly in the western U.S., where recent changes are linked with climate warming and 20th-century land management practices. OBJECTIVES: To study forest resilience to recent wildfires, we examined relationships among fire legacies, landscape features, ecological conditions, and patterns of post-fire conifer regeneration. METHODS: We quantified regeneration across 182 sites in 21 recent large fires in dry mixed-conifer forests of the U.S. northern Rockies. We used logistic and negative binomial regression to predict the probability of establishment and abundance of conifers 5–13 years post-fire. RESULTS: Seedling densities varied widely across all sites (0–127,500 seedlings ha⁻¹) and were best explained by variability in distance to live seed sources (β = −0.014, p = 0.002) and pre-fire tree basal area (β = 0.072, p = 0.008). Beyond 95 m from the nearest live seed source, the probability of seedling establishment was low. Across all the fires we studied, 75 % of the burned area with high tree mortality was within this 95-m threshold, suggesting the presence of live seed trees to facilitate natural regeneration. CONCLUSIONS: Combined with the mix of species present within the burn mosaic, dry mixed-conifer forests will be resilient to large fires across our study region, provided that seedlings survive, fire do not become more frequent, high-severity patches do not get significantly larger, and post-fire climate conditions remain suitable for seedling establishment and survival.

Background Wildfires, like many disturbances, can be catalysts for ecosystem change. Given projected climate change, tree regeneration declines and ecosystem shifts following severe wildfires are predicted. We reviewed scientific literature on post-fire tree regeneration to understand where and why no or few trees established. We wished to distinguish sites that won’t regenerate to trees because of changing climate from sites where trees could grow post fire if they had a seed source or were planted, thus supporting forest ecosystem services for society and nature, such as timber supply, habitat, watershed protection, and carbon storage. Results Our literature review showed that little to no post-fire tree regeneration was more common in low-elevation, dry forest types than in high-elevation forest types. However, depending on the region and species, low tree regeneration was also observed in high elevation, moist forests. Regeneration densities varied by species and seedling densities were attributed to distances to a seed source, water stress or precipitation, elevation, slope, aspect, and plant competition. Our findings provide land managers with two primary considerations to offset low tree regeneration densities. First, we supply a decision support tool of where to plant tree seedling in large high severity burned patches. Second, we recommend possibilities for mitigating and limiting large high severity burned patches to increase survival of trees to be sources of seed for natural regeneration. Conclusions Few or no tree seedlings are establishing on some areas of the 150+ forest fires sampled across western US, suggesting that forests may be replaced by shrublands and grasslands, especially where few seed source trees survived the wildfires. Key information gaps on how species will respond to continued climate change, repeated disturbances, and other site factors following wildfires currently limit our ability to determine future trends in forest regeneration. We provide a decision tree to assist managers in prioritizing post-fire reforestation. We emphasize prioritizing the interior of large burned patches and considering current and future climate in deciding what, when, and where to plant trees. Finally, managing fires and forests for more seed-source tree survival will reduce large, non-forested areas following wildfires where post-fire management may be necessary.

The future of temperate forests in the face of global change and anthropogenic stressors remains uncertain. The regeneration stage, which is a critical bottleneck for many organisms, is a key indicator of forest health, future canopy composition and forest adaptive capacity. In trees, seemingly healthy forests can be at long‐term risk due to insufficient juveniles to replace them (regeneration failure), or compositional differences between juveniles and adults (regeneration mismatch). We propose ‘regeneration debt’ to collectively describe regeneration failure and mismatch in analogy to extinction debt. To demonstrate this concept, we conducted a macroecological analysis of regeneration debt and anthropogenic stressors in eastern US forests. Using U.S. Forest Service‐Forest Inventory and Analysis data, we quantified regeneration debt in 18 states from Maine to South Carolina, and evaluated the influence of site, anthropogenic stressors and climate drivers in the most affected regions. We identified three distinct regions, with little debt in the north, moderate debt in the south and severe regeneration debt in the central, mid‐Atlantic region. In this region, multiple anthropogenic stressors (invasive plants, deer overabundance and land use) were associated with both low‐regeneration abundance and the prevalence of disease‐prone and/or suboptimal species. Synthesis and applications. Without management intervention, the severe regeneration debt in the mid‐Atlantic region will likely lead to long‐term declines in forest cover, with cascading negative effects on forest‐dependent taxa and ecosystem services. Moreover, the location of the regeneration debt, which is at the northern edge of and involves many of the tree species that are predicted to gain suitable habitat in the Northeastern US, has consequences that extend far beyond its current geographic extent. In fact, this regeneration debt may already be functioning as a barrier to poleward tree migration. Our results demonstrate the value of regeneration debt as an indicator of ecosystem health and forest adaptive capacity. Without management intervention, the severe regeneration debt in the mid‐Atlantic region will likely lead to long‐term declines in forest cover, with cascading negative effects on forest‐dependent taxa and ecosystem services. Moreover, the location of the regeneration debt, which is at the northern edge of and involves many of the tree species that are predicted to gain suitable habitat in the Northeastern US, has consequences that extend far beyond its current geographic extent. In fact, this regeneration debt may already be functioning as a barrier to poleward tree migration. Our results demonstrate the value of regeneration debt as an indicator of ecosystem health and forest adaptive capacity.

1. Drought events occurring under warmer temperatures (i.e. \"hotter droughts\") have resulted in widespread tree mortality across the globe, and may result in biomelevel vegetation shifts to alternate vegetation types if there is a failure of trees to regenerate.We investigated how overstorey trees, understorey vegetation, and local climatic and edaphic conditions interact to influence tree regeneration, a key prerequisite for resilience, in a region that has experienced severe overstorey tree mortality due to hotter droughts and beetle infestations. 2. We used detailed field observations from 142 sites that spanned a broad range of environmental conditions to evaluate the effects of climate and recent tree mortality on tree regeneration dynamics in the spatially extensive piñon (Pinus edulis)- juniper (Juniperus osteosperma, Juniperus monosperma) woodland vegetation type of the southwestern USA. We used a structural equation modelling framework to identify how tree mortality and local climatic and edaphic conditions affect piñon and juniper regeneration and electivity analyses to quantify the species-specific associations of tree juveniles with overstorey trees and understorey shrubs. 3. Piñon regeneration appears to be strongly dependent upon advanced regeneration, (i.e. the survival of juvenile trees that established prior to the mortality event), the survival of adult seed-bearing trees (inferred from basal area of surviving trees) and the facilitative effects of overstorey trees for providing favourable microsites for seedling establishment. Model results suggest that local edaphoclimatic conditions directly affected piñon and juniper regeneration, such that stands with hotter, drier local climatic conditions and lower soil available water capacity had limited tree regeneration following large-scale dieback. 4. Synthesis. We identify four indicators of resilience to hotter drought conditions: (1) abundant advance regeneration of tree seedlings; (2) sufficient canopy cover for survival of emergent seedlings and existing regeneration; (3) sufficient seed source from surviving trees with high reproductive output; (4) areas with cooler and wetter local climates and greater soil available water capacity. In the absence of these conditions, there is greater likelihood of woodlands transitioning to more xeric vegetation types following dieback.

Aim To determine if differences in climate envelopes for six coniferous tree species and two life stages (trees and seedlings) suggest a potential for species range contractions, expansions or shifts in response to climate change and if these patterns differ between subalpine (i.e. cool-climate) and montane (i.e. warm-climate) species. Location The dry domain of the western United States. Methods Using data from the Forest Inventory and Analysis National Program, we quantified the relationship between probability of occurrence and climate for adults and seedlings of each species with a Bayesian logistic regression. Assuming that distributional differences between life stages highlight shifting regeneration patterns relative to adult trees, we assessed differences between seedlings and adult trees based on predicted probabilities of occurrence and climate envelope boundaries. Results Differences between occurrence probabilities for seedlings and adults were greatest for montane, as opposed to subalpine, species and along range margins, especially in the southern and western portions of the study area. Climate envelope boundaries of seedlings differed from adult trees most frequently in montane species and often suggested range contractions or range shifts, as opposed to range expansion. Main conclusions Our results indicated that climate-induced contractions and shifts in seedling distribution in response to recent change are already under way and are particularly severe in montane tree species. While adult trees may persist for hundreds of years without significant regeneration, tree species ranges will eventually contract where tree regeneration fails.

Variable retention (VR) has been adopted globally as an alternative to more intensive forms of regeneration harvest. By retaining live trees within harvest units, VR seeks balance among the commodity, ecological, and aesthetic values of managed forests. Achieving these multiple, often competing objectives requires an understanding of how level and spatial pattern of retention shape the abundance, growth, and mortality of regenerating and retained trees. Using long-term (18–19 yr) data from a regional-scale VR experiment, we explore the individual and interactive effects of retention level (15% vs. 40% of initial basal area) and pattern (dispersed vs. aggregated) on the post-harvest dynamics of forests of differing structure and seral composition. Level and pattern of retention imposed trade-offs for the density and growth of regenerating trees (≥0.1 m tall, <5 cm dbh) and ingrowth (trees attaining 5 cm during the study). Greater retention led to greater density of late-seral regeneration, but lower density of early-seral ingrowth, and slower growth of late-seral ingrowth. Dispersed retention enhanced the density of early- and late-seral regeneration (compared to aggregated treatments), but reduced the growth of early-seral ingrowth. We also observed trade-offs for retained trees. Lower retention enhanced the growth of smaller trees (<25 cm dbh), particularly in dispersed settings, but reduced the survival of larger trees, which were more susceptible to windthrow. Greater retention reduced the growth, but enhanced the survival of smaller trees. Pattern imposed similar trade-offs, with dispersed retention enhancing growth, but reducing survival of small trees. Finally, level and pattern resulted in trade-offs for productivity of regenerating vs. retained-tree cohorts. Ingrowth productivity was greater at lower retention and in aggregated treatments; retained-tree productivity was greater at higher retention and in dispersed treatments. Our results provide a unique, long-term perspective on the sensitivity of tree regeneration, growth, and mortality to key structural elements of VR systems. Strong responses to level and pattern of retention produce trade-offs for different ecological or resource objectives. Balancing these objectives may require the combined use of aggregates, dispersed retention, and clearings, to mimic the spatial heterogeneity of habitats, physical structures, and resource conditions that are produced by natural disturbances.